ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2 , Winter 2011

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Numerical of Phalaris () Based on Morphological Characteristics

Maryam Keshavarzi1*, Mahnaz Khaksar 2, Mahvash Seifali 1, Parinaz Ghadam1 1- Department of Biology, Faculty of Science, Alzahra University, Vanak, Tehran, Iran. 2- Graduated M.Sc. Student of systematic, Department of Biology, Faculty of Science, Alzahra University, Vanak, Tehran, Iran.

ﺗﺎﮐﺴﻮﻧﻮﻣﯽ ﻋﺪدي ﮔﻮﻧ ﻪﻫﺎي Phalaris از ﺗﯿﺮه ﻏﻼت ﺑﺮ اﺳﺎس ﺻﻔﺎت رﯾﺨﺖ ﺷﻨﺎﺳﯽ

ﻣﺮﯾﻢ ﮐﺸﺎورزي 1*، ﻣﻬﻨﺎز ﺧﺎﮐﺴﺎر2، ﻣﻬﻮش ﺳﯿﻔﻌﻠﯽ1، ﭘﺮﯾﻨﺎز ﻗﺪم Abstract 1 1- ﮔﺮوه زﯾﺴ ﺖﺷﻨﺎﺳﯽ، داﻧﺸﮑﺪه ﻋﻠﻮم ﭘﺎﯾﻪ، داﻧﺸﮕﺎه اﻟﺰﻫﺮا، وﻧﮏ، ﺗﻬﺮان، Phalaris is a grass which is distributed throughout اﯾﺮان. the temperate regions of the world. There are many 2- داﻧ ﺶآﻣﻮﺧﺘﻪ ﮐﺎرﺷﻨﺎﺳﯽ ارﺷﺪ ﺳﯿﺴﺘﻤﺎﺗﯿﮏ ﮔﯿﺎﻫﯽ ، ﮔﺮوه زﯾﺴ ﺖﺷﻨﺎﺳﯽ، taxonomic problems in this genus. In order to داﻧﺸﮑﺪه ﻋﻠﻮم ﭘﺎﯾﻪ، داﻧﺸﮕﺎه اﻟﺰﻫﺮا، وﻧﮏ، ﺗﻬﺮان، اﯾﺮان. clarify the taxonomy and the interrelationships among Iranian Phalaris taxa, including Ph. minor, ﭼﮑﯿﺪه Ph. arundinaceae, Ph. brachystachys and Ph. ﺟﻨﺲ Phalaris واﺟﺪ ﮔﯿﺎﻫﺎﻧﯽ ﻋﻠﻔﯽ ﺑﺎ ﺗﻮزﯾﻌﯽ ﮔﺴﺘﺮده در ﻣﻨﺎﻃﻖ ﻣﻌﺘﺪﻟﻪ paradoxa 70 qualitative and quantitative دﻧﯿﺎﺳﺖ. ﻣﺸﮑﻼت ﺗﺎﮐﺴـﻮﻧﻮﻣﯿﮑﯽ ﻣﺘﻌـﺪدي در اﯾـﻦ ﺟـﻨﺲ وﺟـﻮد دارد . ﺑـﻪ .morphological characteristics were examined ﻣﻨﻈﻮر روﺷـﻦ ﺳـﺎﺧﺘﻦ وﺿـﻌﯿﺖ ﺗﺎﮐﺴـﻮﻧﻮﻣﯽ و رواﺑـﻂ ﻣﯿـﺎن ﺗﺎﮐﺴـﻮن ﻫـﺎي Numerical taxonomy was performed on 36 Phalaris در اﯾﺮان، ﻣﺸﺘﻤﻞ ﺑﺮ ,accessions of 5 taxa, concerning intra- and inter- Ph. minor, Ph. arundinaceae populations variations as well as inter-specific Ph. brachystachys و Ph. Paradoxa در ﻣﺠﻤﻮع 70 ﺻﻔﺖ ﮐﻤﯽ relationships. The most variable morphological

و ﮐﯿﻔﯽ رﯾﺨﺖﺷﻨﺎﺧﺘﯽ ﻣﻮرد ﺑﺮرﺳﯽ واﻗﻊ ﺷﺪ. ﺗﺎﮐﺴﻮﻧﻮﻣﯽ ﻋـﺪدي ﺑـﺮ روي characters in the species delimitation were also 36 واﺣﺪ ﺟﻤﻌﯿﺘﯽ از 5 ﺗﺎﮐﺴﻮن ﺑﺎ در ﻧﻈﺮ ﮔﺮﻓﺘﻦ ﺗﻨـﻮع درون ﺟﻤﻌﯿﺘـﯽ، ﺑـﯿﻦ determined. The species differed significantly in most of chosen qualitative characters. Results show ﺟﻤﻌﯿﺘﯽ و ﺑﯿﻦ ﮔﻮﻧ ﻪاي اﻧﺠﺎم ﺷﺪ. ﻣﺘﻐﯿ ﺮﺗﺮﯾﻦ ﺻﻔﺎ ترﯾﺨﺘﯽ در ﺗﻌﯿﯿﻦ ﺣـﺪود a clear similarity between two varieties of Ph. ﮔﻮﻧﻪﻫﺎ ﻣﺸﺨﺺ ﺷﺪﻧﺪ. ﮔﻮﻧﻪﻫﺎ ﺗﻔﺎوت ﻣﻌﻨـﯽ داري در ﻏﺎﻟـﺐ ﺻـﻔﺎت ﮐﯿﻔـﯽ paradoxa. Despite the wide range of habitats where اﻧﺘﺨــﺎب ﺷــﺪه ﻧﺸــﺎن دادﻧــﺪ . ﻧﺘــﺎﯾﺞ ﻣﻮﯾــﺪ ﺷــﺒﺎﻫﺖ ﺑــﺎﻻﯾﯽ ﺑــﯿﻦ دو وارﯾﺘــﻪ the accessions were gathered, they showed four Ph. Paradoxa اﺳﺖ. ﻋﻠﯽرﻏﻢ ﺗﻨﻮع زﯾﺎد در زﯾﺴـﺘﮕﺎه ﻫـﺎي واﺣـﺪﻫﺎي definite spots in their PCA ordination graphs. This showed the efficiency of chosen morphological ﺟﻤﻌﯿﺘﯽ ﻣﻮرد ﺑﺮرﺳﯽ،آ نﻫﺎ ﭼﻬﺎر ﺗـﻮده ﻣﺸـﺨﺺ را در ﻧﻤـﻮدار رﺳـﺘﻪ ﺑﻨـﺪي characters in this study. An identification key based ﻣﺸﺨﺺ ﺳﺎﺧﺘﻨﺪ. اﯾﻦ اﻣﺮ ﮐـﺎرآﯾﯽ ﺻـﻔﺎت رﯾﺨﺘـﯽ اﻧﺘﺨـﺎ ب ﺷـﺪه را در اﯾـﻦ on studied features was provided and the species

ﺑﺮرﺳﯽ ﻧﺸﺎن ﻣﯽدﻫﺪ. ﮐﻠﯿﺪ ﺷﻨﺎﺳﺎﯾﯽ ﺑﺮ ﻣﺒﻨﺎي ﺻﻔﺎت ﻣﻮرد ﺑﺮرﺳﯽ اراﺋﻪ ﺷﺪه .relationships were discussed و رواﺑﻂ ﺑﯿﻦ ﮔﻮﻧﻪ ﻫﺎ ﻣﻮرد ﺑﺤﺚ ﻗﺮار ﮔﺮﻓﺘﻪ اﺳﺖ. Keywords: Cluster analysis, Ordination, Phalaris, ﮐﻠﻤﺎت ﮐﻠﯿﺪي: ﺗﺠﺰﯾﻪ ﺧﻮﺷ ﻪاي، رﺳﺘﻪ ﺑﻨﺪي، Phalaris و ﻗﺮا ﺑﺖﻫﺎ. relationship

* Corresponding author. E-mail Address: [email protected] ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 29 ¡ ¡ Introduction morphological characteristics (Tables 2 and 3) selected The genus phalaris L. has had a complicated on the basis of the studies of Bor (1968 & 1970), taxonomic and nomenclatural history (Baldini, 1993 & Baldini (1993 & 1995) and our own field observations. 1995). Phalaris comprises 22 species of annual and For each population, 10 individuals were studied for perennial grasses in temperate regions throughout the their qualitative and quantitative morphological world. It is commonly adventives species of open characteristics. habitats. Phalaris species are distributed throughout In order to detect significant differences in studied various regions of Iran and are among important characteristics among populations of each species and forage of Iran. There are four species and 5 taxa also among different species, analysis of variance of Phalaris in Iran (Bor, 1970): Ph. minor Retz., Ph. (ANOVA) followed by the least significant differences brachystachys Link., Ph. paradoxa L. (with 2 (LSD) tests, were performed. varieties) and Ph. arundinaceae L. Members of the To reveal species relationships cluster analysis and genus Phalaris display many variations on the principal component analysis (PCA) was used standard structure of the inflorescence (Bor, 1968). There is no report of a systematic study on the (Ingrouille, 1986). For multivariate analysis the mean Phalaris species of Iran. The objective of this study of quantitative characteristics was used while was to assess the level of morphological variation of qualitative characters were coded as binary/multi-state 36 accessions of four Phalaris species. We have tried characteristics. Standardized variables used for to reveal the inter-population morphological variations multivariate statistical analysis. The average and inter-specific relationships for the first time. taxonomic distances and squared Euclidean distances

were used as the dissimilarity coefficient in cluster Materials and Methods analysis of morphological data. In order to determine Morphometric studies performed on 36 populations of the most variable morphological characters among the 4 species (5 taxa) native to Iran (Table 1). Voucher studied species, factor analysis based on principal specimens were deposited in the herbarium of Alzahra University. We studied 70 quantitative and qualitative components analysis performed. We used SPSS ver. 9 (1998) software for statistical analysis.

Table 1- Voucher details of Phalaris sampled in this study. Taxon Locality, Height from S.L and collector Herb. No. Ph. minor Yazd, Ardakan, 1400 m, Keshavarzi. 83m8 Khozestan, Masjed Soleiman, Boromandi. 68m12 Tehran, Chitgar, Keshavarzi. 80m16 Yazd, Meyboud, 1400 m, Keshavarzi. 83m14 Tehran, Saeii Park, 1600 m, Keshavarzi. 66m22 Qoum, Vejdani. 71m17 Markazi, Mahalat, Vejdani. 70m18 Khozestan, Ramhormouz, Keshavarzi. 83m18 Fars, Shiraz, 1500 m, Keshavarzi. 84m10 Khozestan, Molasani, Farasat. 83m5 Markazi, Saveh, Yalabad, Keshavarzi. 83m3 Lorestan, Khoramabad, Kew Park, Keshavarzi. 83m7 Khozestan, Hafttapeh, Choghazanbil, Keshavarzi. 83m13 Khozestan, Behbahan, Keshavarzi. 83m6 Tehran, Vanak 1700 m, Keshavarzi. 83m19 Mazandaran, Sari, Komeilidost. 82m15 Fars, Kazeroun, Baghenazar, 900 m , Keshavarzi. 84m1 Khozestan, Ahvaz, Keshavarzi. 83m4 Golestan, Gorgan, Khaksar. 83m23

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 30 ¡ ¡

Continue Table 1 Taxon Locality, Height from S.L and collector Herb. No. Ph. brachystachys Mazandaran, Sari, Dashtenaz, Khaksar 1385b2 Khozestan, Ramhormouz, Bayman arayez Village, Farasat. 1385b1 Mazandaran, Sari, Sameskandeh Khaksar. 1385b3 Tehran, Khaksar. 1366b6 Khozestan, Izae, Eshkafte Soleiman, Keshavarzi. 1383b7 Khozestan, Molasani, Farasat. 138b4 Tehran, Karaj, Mardabad, Keshavarzi. 1383b8 Ph. paradoxa var. paradoxa Khozestan, Molasani, Farasat. 85pp1 Khozestan, 15 km to Izae, Nanaii. 85pp3 Khozestan, Ramhormouz, Bayman Arayez Village, Farasat. 85pp2 Ph. paradoxa var. Mazandaran, Sari, Bayekola, 15 km Nesa, Khaksar. 85pps1 praemorsa Mazandaran, Savadkooh, zir Abe joush Village, Khaksar. 85pps1 Mazandaran, Sari, Sameskandeh, Khaksar. 85pp7 Golestan, Gorgan, Khaksar. 85pps6 Mazandaran, Behshahr, Khaksar. 85pps9 Mazandaran, Dashtenaz, Khaksar. 85pps5 Ph. arundinaceae Tehran to Chalous, Dizin, Khaksar. 85a1

Table 2 - Quantitative morphological characteristics in the Phalaris species studied. All scales are in mm. Character Character Plant height Length of lowermost Glume Palae width Width of lowermost Glume Length of longest internode Length of Lemma Caryopsis length Width of Lemma Caryopsis width Length of palae Leaf number Length/width of caryopsis Leaf width Plant height/length of longest internode Leaf length Length/width of leaf blade Ligule Length Length/width of spike Spikelet Length Length /width of spikelet Spikelet width Length/width of lowermost glume Spike length Length/width of uppermost Glume Spike width Length/width of lemma Floret number Length/width of palae Length of uppermost Glume Length of spike to length of spikelet Width of uppermost Glume Width of Spike to Width of spikelet

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 31 ¡ ¡ T able 3 - Qualitative morphological characteristics In the Phalaris species studied. No Character State of Character 1 Plant habit Annual (0), Perennial (1) 2 Culms direction Erect(0), Curved (1) 3 Inflorescence compactness Dense(0), Lax (1) 4 Fertility of Spikelets All fertile (0), Fertile and Sterile (1) 5 Inflorescence Rachis Fragile (0), Tough (1) 6 Spike Shape Ovoid(0), elliptic (1), Oblanceolate (2) 7 Spike covering Dense (0), sparse (1) 8 Spike compactness Compact (0), Lax(1) 9 Leaf surface Smooth (0), Rough (1) 10 Leaf Color Pale Green (0), Dark Green (1) 11 Leaf sheath Covering spike(0), Not so (1) 12 Shape of Ligule apex Truncate (0), Rounded (1), Fringed (2), Cylindrical(3) 13 Ligule covering Smooth(0), Hairy (1) 14 Spikelet Surface Hairy (0), Smooth (1) 15 Sterile Spikelets Uniform (0), Some club like (1) 16 Rudimentary spikelets Present (0), Absent (1) 17 Glume Color Pale Green (0), Dark Green (1) 18 Glume Apex Longley pointed (0), Shortly pointed (1) 19 Glume Covering Smooth(0), Hairy (1) 20 Glume Hair Density Dense (0), sparse (1) 21 Glume nerves Convergent (0), Not so (1) 22 Winged Glume Present (0), Absent (1) 23 Glume Wing Tooth like (0), Not so (1) 24 Glume Wing Margins Smooth (0), Undulate (1) 25 Glume Wing Texture Membranous(0), non-membranous (1) 26 Shape of lemma Apex Pointed (0), rounded (1) 27 Lemma Texture Membranous(0), non-membranous (1) 28 Lemma shape Ovoid (0), elliptic – oblong (1) 29 Lemma nerves Present (0), absent (1) 30 Lemma Nerves Convergent (0), not so (1) 31 Lemma Hairs Dense (0), sparse (1) 32 Lemma hairs Length Long (0), Short (1) 33 Palae Texture Nerved (0), not so (1) 34 Palae hairs Present (0), Absent (1) 35 Palae hair position At margins (0), distributed (1) 36 Palae hairs Length Long (0), Short (1) 37 Caryopsis color Brown (0), Yellow (1), Green (2) 38 Caryopsis Shape Ovoid (0), Elliptic (1), Lanceolate (2)

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 32 ¡ ¡ Results and qualitative characteristics produced similar results The analyses showed that the range of quantitative (Fig 2). In the overall morphological analysis, two characters is somehow different from what is given in major clusters are formed. The first major cluster is different flora and species descriptions. The species comprised of two sub-clusters, in the first of which studied also differed significantly in most of the populations belonging to Ph. brachystachys. The other quantitative characteristics as revealed by ANOVA sub-cluster is comprised of two sub-clusters itself, one of which contains Ph. minor while Ph. paradoxa is and LSD tests. The mean of such quantitative placed in the other. Ph. arundinaceae is placed in a characters may be useful for species delimitation. This separate cluster. Thus it is evident Ph. minor and Ph. is supported by the clustering (Fig 1.) of the studied paradoxa are closely related which, in turn, are both species based on merely quantitative morphological related to Ph. brachystachys. Ph. minor and Ph. characteristics. Figure 1 show that Ph. arundinaceae arunidinaceae were considered to be closely related to makes a separate cluster and populations of Ph. each other based on having merely fertile spikelets in brachystachys make separate ones. their inflorescences in the Flora Iranica (Bor, 1970). The cluster analysis and PCA ordination of He also considered a separate place for Ph. Phalaris species of Iran, based on both quantitative arundinaceae due to its perennial habit.

Figure 1- WARD clustering of Phalaris species based on the mean of quantitative morphological characters.

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 33 ¡ ¡ In order to determine the most variable characters or geographical forms. The use of similar methods in among the studied species, factor analysis based on this present study has indicated that, two varieties of PCA was performed revealing that the first two factors Ph. paradoxa (praemorsa and Paradoxa) make a comprise about 77% of total variation. In the first sister group (Fig. 1). These two show the nearest factor with about 58% of total variation (Table 4), relationship with Ph. minor. Ph. brachystachys in characteristics such as spike width, length of first level 10 and show a relationship with these clusters. inter-node and spikelet, width of leaf blade and leaf Ph. arundinaceae which is the only perennial species number, leaf number, spike length, width of upper in this genus of Iran, forms a separate cluster (level glume, length and width of lemma, width and length 25). Cluster analysis of the anatomical features of of lower glume possessed the highest positive dorsal epidermis and leaf trans-sections by the WARD correlation (≥0.7). method provide further evidence for such a relationship (Keshavarzi et al., 2009). Table 4 - Factor analysis results based on the morphological Considering the results of factor analysis here we characteristics of Phalaris populations of Iran. provide an identification key for Iranian species of Character Factor 1 Factor 2 Phalaris: Spike Width 0.97 1a- Plant annual, Spike compact, glume winged 2 Length of first internode 0.96 b- Plant perennial, spike loose, glume without wing Length of spikelet 0.96 Ph.arundinaceae 2a - All spikelets fertile, glumes' wing dentate, ligule fringed and wide Width of leaf blade 0.95 Ph.minor Leaf number 0.95 b-Spikelets fertile and sterile, glumes' wing not dentate, ligule Spike length 0.95 tip rounded or truncate 3 3a- Rudimentary spikelets present, glumes' wing tooth like 4 Width of upper glume 0.95 b-Rudimentary spikelets absent, glumes' wing complete Length of lemma 0.93 Ph.brachystahys Width of lower glume 0.91 4a- Rudimentary spikelets club like, ligule surface without hair Ph. paradoxa var. praemorsa Lemma width 0.90 b- Rudimentary spikelets not club shaped, ligule surface hairy Length of lower glume 0.72 Ph. paradoxa var. paradoxa Length of upper glume 0.85 Caryopsis width 0.77 Discussion Caryopsis length 0.72 The most polymorphic species in Iran is P. minor. Our results are in accordance with those of Hucle and In the second factor with about 19.4% of total Matus (1999). Their vast variation could be related to variation, characteristics such as the length of the the auto-tetraploid condition of this species. There also upper glume and caryopsis width and length possessed some edaphic conditions which are able to affect ph. the highest positive correlations. Therefore, these are minor populations in different places. We found that the most variable morphological characters among individual size is related to soil conditions. In rich Phalaris species of Iran. soils of fields Ph. minor could reach up to 100 cm and In assessing the inter-specific affinities and intra- bear up to 6 cm spikes, while in poor soils Ph. minor specific variability of a complex group of species, the individuals will be 10-15 cm height and bear only 1.5 multivariate analysis methods had been shown to be cm spikes (Keshavarzi et al., 2007 b). very helpful. Such methods were used by Ng et al. Anderson (1961) clarified the genetic relationships (1981) in their studies of rice species to clarify the of annual species of phalaris. He believed that four inter-specific relationships and to distinguish species types of variations occurred in sterile florets of this

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 34 ¡ ¡

Figure 2- PCA ordination of the Phalaris species b ased on morphological characteristics.

genus. He defined four definite lineages in this genus: References Ph. paradoxa, Ph. brachystachys, Ph. arundinaceae Anderson, D.E. (1961). Taxonomy and distribution of and ten other species containing Ph. minor. He the genus Phalaris. Iowa state Journal of Science, separated the last group by their geographical 36:1-96. distribution pattern to old world (eg: Ph. minor) and new world. Baldini (1995) believed that there are two Baldini, R.M. (1995). Revision of the genus Phalaris centers of diversification for these taxa, one of which L. (Gramineae). Webbia, 49:265-329. is Mediterranean area and the other is south western USA. Iranian species of Phalaris are of the old world Baldini, R.M. (1993). The genus Phalaris L. group. Baldini believed that Ph. arundinaceae is the (Gramineae) in Italy. Webbia, 47: 1-53. only species that connects new and old world species. As the Phalaris migration from the Mediterranean area Bor, N.L. (1968). Gramineae. In: Townsend, C., E. occurred during the tertiary period, we conclude that Guest. and Al-Rawi, A., (eds.), Flora of Iraq. Phalaris species of Iran have experienced a relatively 9:361-370. Baghdad: Iraq Ministry of long period of divergence and adaptations to local Agriculture. conditions in their habitats in Iran.

ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 35 ¡ ¡ Bor, N.L. (1970). Gramineae. In: Rechinger, K. H. Ng, N.Q., T.T. Chang, J.T. Williams and J.G. Hawkes (ed.) Flora Iranica. 70. 345-349, Graz. Austria: (1981). Morphological studies of Asian rice and Akademische Druk - Und Verlasantalt.Wiena. its related wild species and the recognition of a new Australian taxon. Biological Journal of Hucle, P.G. and M. Matus (1999). Isozyme Variation Linnean Society, 16: 303-313. within and among Accessions of annual Phalaris species in North American Germplasm Collection. Crop Science, 39: 1222-1228.

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ﻋـﻠـﻮم ﻣﺤـﯿـﻄﯽ ﺳﺎل ﻫﺸﺘﻢ، ﺷﻤﺎره دوم، زﻣﺴﺘﺎن 1389 ENVIRONMENTAL SCIENCES Vol.8, No.2, Winter 2011 36 ¡ ¡